Chris Field

Melvin and Joan Lane Professor for Interdisciplinary Environmental Studies, Director, Woods Institute for the Environment, Professor of Earth System Science, of Biology and Senior Fellow at the Precourt Institute for Energy

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ResearchMy field is global ecology, and my research emphasizes ecological contributions across the range of Earth science disciplines. My colleagues and I develop diverse approaches to quantifying large-scale ecosystem processes, using satellites, atmospheric data, models, and census data, and explore global-scale patterns of vegetation-climate feedbacks, carbon cycle dynamics, primary production, forest management, and fire. At the ecosystem-scale, we conduct experiments on grassland responses to global change, which integrate approaches from molecular biology to remote sensing.

TeachingI am one of five professors who teach the Earth Systems field studies course for advanced undergrads and co-terms at Jasper Ridge Biological Preserve. I also teach an introductory seminar on climate change for freshmen.

Abstract

Understanding the pathways through which drought stress kills woody vegetation can improve projections of the impacts of climate change on ecosystems and carbon-cycle feedbacks. Continuous in situ measurements of whole trees during drought and as trees die hold promise to illuminate physiological pathways but are relatively rare. We monitored leaf characteristics, water use efficiency, water potentials, branch hydraulic conductivity, soil moisture, meteorological variables, and sap flux on mature healthy and sudden aspen decline-affected (SAD) trembling aspen (Populus tremuloides) ramets over two growing seasons, including a severe summer drought. We calculated daily estimates of whole-ramet hydraulic conductance and modeled whole-ramet assimilation. Healthy ramets experienced rapid declines of whole-ramet conductance during the severe drought, providing an analog for what likely occurred during the previous drought that induced SAD. Even in wetter periods, SAD-affected ramets exhibited fivefold lower whole-ramet hydraulic conductance and sevenfold lower assimilation than counterpart healthy ramets, mediated by changes in leaf area, water use efficiency, and embolism. Extant differences between healthy and SAD ramets reveal that ongoing multi-year forest die-off is primarily driven by loss of whole-ramet hydraulic capability, which in turn limits assimilation capacity. Branch-level measurements largely captured whole-plant hydraulic limitations during drought and mortality, but whole-plant measurements revealed a potential role of other losses in the hydraulic continuum. Our results highlight the importance of a whole-tree perspective in assessing physiological pathways to tree mortality and indicate that the effects of mortality on these forests' assimilation and productivity are larger than expected based on canopy leaf area differences.

Abstract

Solar energy installations in deserts are on the rise, fueled by technological advances and policy changes. Deserts, with a combination of high solar radiation and availability of large areas unusable for crop production are ideal locations for large solar installations. However, for efficient power generation, solar infrastructures use large amounts of water for construction and operation. We investigated the water use and greenhouse gas (GHG) emissions associated with solar installations in North American deserts in comparison to agave-based biofuel production, another widely promoted potential energy source from arid systems. We determined the uncertainty in our analysis by a Monte Carlo approach that varied the most important parameters, as determined by sensitivity analysis. We considered the uncertainty in our estimates as a result of variations in the number of solar modules ha(-1), module efficiency, number of agave plants ha(-1), and overall sugar conversion efficiency for agave. Further, we considered the uncertainty in revenue and returns as a result of variations in the wholesale price of electricity and installation cost of solar photovoltaic (PV), wholesale price of agave ethanol, and cost of agave cultivation and ethanol processing. The life-cycle analyses show that energy outputs and GHG offsets from solar PV systems, mean energy output of 2405 GJ ha(-1) year(-1) (5 and 95% quantile values of 1940-2920) and mean GHG offsets of 464 Mg of CO2 equiv ha(-1) year(-1) (375-562), are much larger than agave, mean energy output from 206 (171-243) to 61 (50-71) GJ ha(-1) year(-1) and mean GHG offsets from 18 (14-22) to 4.6 (3.7-5.5) Mg of CO2 equiv ha(-1) year(-1), depending upon the yield scenario of agave. Importantly though, water inputs for cleaning solar panels and dust suppression are similar to amounts required for annual agave growth, suggesting the possibility of integrating the two systems to maximize the efficiency of land and water use to produce both electricity and liquid fuel. A life-cycle analysis of a hypothetical colocation indicated higher returns per m(3) of water used than either system alone. Water requirements for energy production were 0.22 L MJ(-1) (0.28-0.19) and 0.42 L MJ(-1) (0.52-0.35) for solar PV-agave (baseline yield) and solar PV-agave (high yield), respectively. Even though colocation may not be practical in all locations, in some water-limited areas, colocated solar PV-agave systems may provide attractive economic incentives in addition to efficient land and water use.

Abstract

Secondary forests cover large areas of the tropics and play an important role in the global carbon cycle. During secondary forest succession, simultaneous changes occur among stand structural attributes, soil properties, and species composition. Most studies classify tree species into categories based on their regeneration requirements. We use a high-resolution secondary forest chronosequence to assign trees to a continuous gradient in species successional status assigned according to their distribution across the chronosequence. Species successional status, not stand age or differences in stand structure or soil properties, was found to be the best predictor of leaf trait variation. Foliar δ(13)C had a significant positive relationship with species successional status, indicating changes in foliar physiology related to growth and competitive strategy, but was not correlated with stand age, whereas soil δ(13)C dynamics were largely constrained by plant species composition. Foliar δ(15)N had a significant negative correlation with both stand age and species successional status, - most likely resulting from a large initial biomass-burning enrichment in soil (15)N and (13)C and not closure of the nitrogen cycle. Foliar %C was neither correlated with stand age nor species successional status but was found to display significant phylogenetic signal. Results from this study are relevant to understanding the dynamics of tree species growth and competition during forest succession and highlight possibilities of, and potentially confounding signals affecting, the utility of leaf traits to understand community and species dynamics during secondary forest succession.

Environmental and community controls on plant canopy chemistry in a Mediterranean-type ecosystemPROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICADahlin, K. M., Asner, G. P., Field, C. B.2013; 110 (17): 6895-6900

Abstract

Understanding how and why plant communities vary across space has long been a goal of ecology, yet parsing the relative importance of different influences has remained a challenge. Species-specific models are not generalizable, whereas broad plant functional type models lack important detail. Here we consider plant trait patterns at the local scale and ask whether plant chemical traits are more closely linked to environmental gradients or to changes in species composition. We used the visible-to-shortwave infrared (VSWIR) spectrometer of the Carnegie Airborne Observatory to develop maps of four plant chemical traits-leaf nitrogen per mass, leaf carbon per mass, leaf water concentration, and canopy water content-across a diverse Mediterranean-type ecosystem (Jasper Ridge Biological Preserve, CA). For all four traits, plant community alone was the strongest predictor of trait variation (explaining 46-61% of the heterogeneity), whereas environmental gradients accounted for just one fourth of the variation in the traits. This result emphasizes the critical role that species composition plays in mediating nutrient and carbon cycling within and among different communities. Environmental filtering and limits to similarity can act strongly, simultaneously, in a spatially heterogeneous environment, but the local-scale environmental gradients alone cannot account for the variation across this landscape.

Abstract

Forest mortality constitutes a major uncertainty in projections of climate impacts on terrestrial ecosystems and carbon-cycle feedbacks. Recent drought-induced, widespread forest die-offs highlight that climate change could accelerate forest mortality with its diverse and potentially severe consequences for the global carbon cycle, ecosystem services, and biodiversity. How trees die during drought over multiple years remains largely unknown and precludes mechanistic modeling and prediction of forest die-off with climate change. Here, we examine the physiological basis of a recent multiyear widespread die-off of trembling aspen (Populus tremuloides) across much of western North America. Using observations from both native trees while they are dying and a rainfall exclusion experiment on mature trees, we measure hydraulic performance over multiple seasons and years and assess pathways of accumulated hydraulic damage. We test whether accumulated hydraulic damage can predict the probability of tree survival over 2 years. We find that hydraulic damage persisted and increased in dying trees over multiple years and exhibited few signs of repair. This accumulated hydraulic deterioration is largely mediated by increased vulnerability to cavitation, a process known as cavitation fatigue. Furthermore, this hydraulic damage predicts the probability of interyear stem mortality. Contrary to the expectation that surviving trees have weathered severe drought, the hydraulic deterioration demonstrated here reveals that surviving regions of these forests are actually more vulnerable to future droughts due to accumulated xylem damage. As the most widespread tree species in North America, increasing vulnerability to drought in these forests has important ramifications for ecosystem stability, biodiversity, and ecosystem carbon balance. Our results provide a foundation for incorporating accumulated drought impacts into climate-vegetation models. Finally, our findings highlight the critical role of drought stress accumulation and repair of stress-induced damage for avoiding plant mortality, presenting a dynamic and contingent framework for drought impacts on forest ecosystems.

Fostering advances in interdisciplinary climate sciencePROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICAShaman, J., Solomon, S., Colwell, R. R., Field, C. B.2013; 110: 3653-3656

Abstract

Tree death from drought and heat stress is a critical and uncertain component in forest ecosystem responses to a changing climate. Recent research has illuminated how tree mortality is a complex cascade of changes involving interconnected plant systems over multiple timescales. Explicit consideration of the definitions, dynamics, and temporal and biological scales of tree mortality research can guide experimental and modeling approaches. In this review, we draw on the medical literature concerning human death to propose a water resource-based approach to tree mortality that considers the tree as a complex organism with a distinct growth strategy. This approach provides insight into mortality mechanisms at the tree and landscape scales and presents promising avenues into modeling tree death from drought and temperature stress.

The roles of hydraulic and carbon stress in a widespread climate-induced forest die-offPROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICAAnderegg, W. R., Berry, J. A., Smith, D. D., Sperry, J. S., Anderegg, L. D., Field, C. B.2012; 109 (1): 233-237

Abstract

Forest ecosystems store approximately 45% of the carbon found in terrestrial ecosystems, but they are sensitive to climate-induced dieback. Forest die-off constitutes a large uncertainty in projections of climate impacts on terrestrial ecosystems, climate-ecosystem interactions, and carbon-cycle feedbacks. Current understanding of the physiological mechanisms mediating climate-induced forest mortality limits the ability to model or project these threshold events. We report here a direct and in situ study of the mechanisms underlying recent widespread and climate-induced trembling aspen (Populus tremuloides) forest mortality in western North America. We find substantial evidence of hydraulic failure of roots and branches linked to landscape patterns of canopy and root mortality in this species. On the contrary, we find no evidence that drought stress led to depletion of carbohydrate reserves. Our results illuminate proximate mechanisms underpinning recent aspen forest mortality and provide guidance for understanding and projecting forest die-offs under climate change.

Abstract

Aboveground biomass (AGB) reflects multiple and often undetermined ecological and land-use processes, yet detailed landscape-level studies of AGB are uncommon due to the difficulty in making consistent measurements at ecologically relevant scales. Working in a protected mediterranean-type landscape (Jasper Ridge Biological Preserve, California, USA), we combined field measurements with remotely sensed data from the Carnegie Airborne Observatory's light detection and ranging (lidar) system to create a detailed AGB map. We then developed a predictive model using a maximum of 56 explanatory variables derived from geologic and historic-ownership maps, a digital elevation model, and geographic coordinates to evaluate possible controls over currently observed AGB patterns. We tested both ordinary least-squares regression (OLS) and autoregressive approaches. OLS explained 44% of the variation in AGB, and simultaneous autoregression with a 100-m neighborhood improved the fit to an r2 = 0.72, while reducing the number of significant predictor variables from 27 variables in the OLS model to 11 variables in the autoregressive model. We also compared the results from these approaches to a more typical field-derived data set; we randomly sampled 5% of the data 1000 times and used the same OLS approach each time. Environmental filters including incident solar radiation, substrate type, and topographic position were significant predictors of AGB in all models. Past ownership was a minor but significant predictor, despite the long history of conservation at the site. The weak predictive power of these environmental variables, and the significant improvement when spatial autocorrelation was incorporated, highlight the importance of land-use history, disturbance regime, and population dynamics as controllers of AGB.

Abstract

Global environmental changes are altering interactions among plant species, sometimes favoring invasive species. Here, we examine how a suite of five environmental factors, singly and in combination, can affect the success of a highly invasive plant. We introduced Centaurea solstitialis L. (yellow starthistle), which is considered by many to be California's most troublesome wildland weed, to grassland plots in the San Francisco Bay Area. These plots experienced ambient or elevated levels of warming, atmospheric CO2, precipitation, and nitrate deposition, and an accidental fire in the previous year created an additional treatment. Centaurea grew more than six times larger in response to elevated CO2, and, outside of the burned area, grew more than three times larger in response to nitrate deposition. In contrast, resident plants in the community responded less strongly (or did not respond) to these treatments. Interactive effects among treatments were rarely significant. Results from a parallel mesocosm experiment, while less dramatic, supported the pattern of results observed in the field. Taken together, our results suggest that ongoing environmental changes may dramatically increase Centaurea's prevalence in western North America.

Abstract

To develop a scheme for partitioning the products of photosynthesis toward different biomass components in land-surface models, a database on component mass and net primary productivity (NPP), collected from FLUXNET sites, was examined to determine allometric patterns of allocation. We found that NPP per individual of foliage (Gfol), stem and branches (Gstem), coarse roots (Gcroot) and fine roots (Gfroot) in individual trees is largely explained (r2 = 67-91%) by the magnitude of total NPP per individual (G). Gfol scales with G isometrically, meaning it is a fixed fraction of G ( 25%). Root-shoot trade-offs were manifest as a slow decline in Gfroot, as a fraction of G, from 50% to 25% as stands increased in biomass, with Gstem and Gcroot increasing as a consequence. These results indicate that a functional trade-off between aboveground and belowground allocation is essentially captured by variations in G, which itself is largely governed by stand biomass and only secondarily by site-specific resource availability. We argue that forests are characterized by strong competition for light, observed as a race for individual trees to ascend by increasing partitioning toward wood, rather than by growing more leaves, and that this competition stronglyconstrains the allocational plasticity that trees may be capable of. The residual variation in partitioning was not related to climatic or edaphic factors, nor did plots with nutrient or water additions show a pattern of partitioning distinct from that predicted by G alone. These findings leverage short-term process studies of the terrestrial carbon cycle to improve decade-scale predictions of biomass accumulation in forests. An algorithm for calculating partitioning in land-surface models is presented.

Abstract

Little is known about the combined impacts of global environmental changes and ecological disturbances on ecosystem functioning, even though such combined impacts might play critical roles in shaping ecosystem processes that can in turn feed back to climate change, such as soil emissions of greenhouse gases.We took advantage of an accidental, low-severity wildfire that burned part of a long-term global change experiment to investigate the interactive effects of a fire disturbance and increases in CO(2) concentration, precipitation and nitrogen supply on soil nitrous oxide (N(2)O) emissions in a grassland ecosystem. We examined the responses of soil N(2)O emissions, as well as the responses of the two main microbial processes contributing to soil N(2)O production--nitrification and denitrification--and of their main drivers. We show that the fire disturbance greatly increased soil N(2)O emissions over a three-year period, and that elevated CO(2) and enhanced nitrogen supply amplified fire effects on soil N(2)O emissions: emissions increased by a factor of two with fire alone and by a factor of six under the combined influence of fire, elevated CO(2) and nitrogen. We also provide evidence that this response was caused by increased microbial denitrification, resulting from increased soil moisture and soil carbon and nitrogen availability in the burned and fertilized plots.Our results indicate that the combined effects of fire and global environmental changes can exceed their effects in isolation, thereby creating unexpected feedbacks to soil greenhouse gas emissions. These findings highlight the need to further explore the impacts of ecological disturbances on ecosystem functioning in the context of global change if we wish to be able to model future soil greenhouse gas emissions with greater confidence.

Direct climate effects of perennial bioenergy crops in the United StatesPROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICAGeorgescu, M., Lobell, D. B., Field, C. B.2011; 108 (11): 4307-4312

Abstract

Biomass-derived energy offers the potential to increase energy security while mitigating anthropogenic climate change, but a successful path toward increased production requires a thorough accounting of costs and benefits. Until recently, the efficacy of biomass-derived energy has focused primarily on biogeochemical consequences. Here we show that the biogeophysical effects that result from hypothetical conversion of annual to perennial bioenergy crops across the central United States impart a significant local to regional cooling with considerable implications for the reservoir of stored soil water. This cooling effect is related mainly to local increases in transpiration, but also to higher albedo. The reduction in radiative forcing from albedo alone is equivalent to a carbon emissions reduction of , which is six times larger than the annual biogeochemical effects that arise from offsetting fossil fuel use. Thus, in the near-term, the biogeophysical effects are an important aspect of climate impacts of biofuels, even at the global scale. Locally, the simulated cooling is sufficiently large to partially offset projected warming due to increasing greenhouse gases over the next few decades. These results demonstrate that a thorough evaluation of costs and benefits of bioenergy-related land-use change must include potential impacts on the surface energy and water balance to comprehensively address important concerns for local, regional, and global climate change.

Agricultural net primary production in relation to that liberated by the extinction of Pleistocene mega-herbivores: an estimate of agricultural carrying capacity?ENVIRONMENTAL RESEARCH LETTERSDoughty, C. E., Field, C. B.2010; 5 (4)

Abstract

The ranges of plants and animals are moving in response to recent changes in climate. As temperatures rise, ecosystems with 'nowhere to go', such as mountains, are considered to be more threatened. However, species survival may depend as much on keeping pace with moving climates as the climate's ultimate persistence. Here we present a new index of the velocity of temperature change (km yr(-1)), derived from spatial gradients ( degrees C km(-1)) and multimodel ensemble forecasts of rates of temperature increase ( degrees C yr(-1)) in the twenty-first century. This index represents the instantaneous local velocity along Earth's surface needed to maintain constant temperatures, and has a global mean of 0.42 km yr(-1) (A1B emission scenario). Owing to topographic effects, the velocity of temperature change is lowest in mountainous biomes such as tropical and subtropical coniferous forests (0.08 km yr(-1)), temperate coniferous forest, and montane grasslands. Velocities are highest in flooded grasslands (1.26 km yr(-1)), mangroves and deserts. High velocities suggest that the climates of only 8% of global protected areas have residence times exceeding 100 years. Small protected areas exacerbate the problem in Mediterranean-type and temperate coniferous forest biomes. Large protected areas may mitigate the problem in desert biomes. These results indicate management strategies for minimizing biodiversity loss from climate change. Montane landscapes may effectively shelter many species into the next century. Elsewhere, reduced emissions, a much expanded network of protected areas, or efforts to increase species movement may be necessary.

Abstract

The quantity of land available to grow biofuel crops without affecting food prices or greenhouse gas (GHG) emissions from land conversion is limited. Therefore, bioenergy should maximize land-use efficiency when addressing transportation and climate change goals. Biomass could power either internal combustion or electric vehicles, but the relative land-use efficiency of these two energy pathways is not well quantified. Here, we show that bioelectricity outperforms ethanol across a range of feedstocks, conversion technologies, and vehicle classes. Bioelectricity produces an average of 81% more transportation kilometers and 108% more emissions offsets per unit area of cropland than does cellulosic ethanol. These results suggest that alternative bioenergy pathways have large differences in how efficiently they use the available land to achieve transportation and climate goals.

Abstract

Converting forest lands into bioenergy agriculture could accelerate climate change by emitting carbon stored in forests, while converting food agriculture lands into bioenergy agriculture could threaten food security. Both problems are potentially avoided by using abandoned agriculture lands for bioenergy agriculture. Here we show the global potential for bioenergy on abandoned agriculture lands to be less than 8% of current primary energy demand, based on historical land use data, satellite-derived land cover data, and global ecosystem modeling. The estimated global area of abandoned agriculture is 385-472 million hectares, or 66-110% of the areas reported in previous preliminary assessments. The area-weighted mean production of above-ground biomass is 4.3 tons ha(-1) y(-1), in contrast to estimates of up to 10 tons ha(-1) y(-1) in previous assessments. The energy content of potential biomass grown on 100% of abandoned agriculture lands is less than 10% of primary energy demand for most nations in North America, Europe, and Asia, but it represents many times the energy demand in some African nations where grasslands are relatively productive and current energy demand is low.

Abstract

Dinitrogen (N(2)) fixation is widely recognized as an important process in controlling ecosystem responses to global environmental change, both today and in the past; however, significant discrepancies exist between theory and observations of patterns of N(2) fixation across major sectors of the land biosphere. A question remains as to why symbiotic N(2)-fixing plants are more abundant in vast areas of the tropics than in many of the mature forests that seem to be nitrogen-limited in the temperate and boreal zones. Here we present a unifying framework for terrestrial N(2) fixation that can explain the geographic occurrence of N(2) fixers across diverse biomes and at the global scale. By examining trade-offs inherent in plant carbon, nitrogen and phosphorus capture, we find a clear advantage to symbiotic N(2) fixers in phosphorus-limited tropical savannas and lowland tropical forests. The ability of N(2) fixers to invest nitrogen into phosphorus acquisition seems vital to sustained N(2) fixation in phosphorus-limited tropical ecosystems. In contrast, modern-day temperatures seem to constrain N(2) fixation rates and N(2)-fixing species from mature forests in the high latitudes. We propose that an analysis that couples biogeochemical cycling and biophysical mechanisms is sufficient to explain the principal geographical patterns of symbiotic N(2) fixation on land, thus providing a basis for predicting the response of nutrient-limited ecosystems to climate change and increasing atmospheric CO(2).

Abstract

Increased production of biomass for energy has the potential to offset substantial use of fossil fuels, but it also has the potential to threaten conservation areas, pollute water resources and decrease food security. The net effect of biomass energy agriculture on climate could be either cooling or warming, depending on the crop, the technology for converting biomass into useable energy, and the difference in carbon stocks and reflectance of solar radiation between the biomass crop and the pre-existing vegetation. The area with the greatest potential for yielding biomass energy that reduces net warming and avoids competition with food production is land that was previously used for agriculture or pasture but that has been abandoned and not converted to forest or urban areas. At the global scale, potential above-ground plant growth on these abandoned lands has an energy content representing approximately 5% of world primary energy consumption in 2006. The global potential for biomass energy production is large in absolute terms, but it is not enough to replace more than a few percent of current fossil fuel usage. Increasing biomass energy production beyond this level would probably reduce food security and exacerbate forcing of climate change.

Abstract

The growth rate of atmospheric carbon dioxide (CO(2)), the largest human contributor to human-induced climate change, is increasing rapidly. Three processes contribute to this rapid increase. Two of these processes concern emissions. Recent growth of the world economy combined with an increase in its carbon intensity have led to rapid growth in fossil fuel CO(2) emissions since 2000: comparing the 1990s with 2000-2006, the emissions growth rate increased from 1.3% to 3.3% y(-1). The third process is indicated by increasing evidence (P = 0.89) for a long-term (50-year) increase in the airborne fraction (AF) of CO(2) emissions, implying a decline in the efficiency of CO(2) sinks on land and oceans in absorbing anthropogenic emissions. Since 2000, the contributions of these three factors to the increase in the atmospheric CO(2) growth rate have been approximately 65 +/- 16% from increasing global economic activity, 17 +/- 6% from the increasing carbon intensity of the global economy, and 18 +/- 15% from the increase in AF. An increasing AF is consistent with results of climate-carbon cycle models, but the magnitude of the observed signal appears larger than that estimated by models. All of these changes characterize a carbon cycle that is generating stronger-than-expected and sooner-than-expected climate forcing.

Abstract

CO2 emissions from fossil-fuel burning and industrial processes have been accelerating at a global scale, with their growth rate increasing from 1.1% y(-1) for 1990-1999 to >3% y(-1) for 2000-2004. The emissions growth rate since 2000 was greater than for the most fossil-fuel intensive of the Intergovernmental Panel on Climate Change emissions scenarios developed in the late 1990s. Global emissions growth since 2000 was driven by a cessation or reversal of earlier declining trends in the energy intensity of gross domestic product (GDP) (energy/GDP) and the carbon intensity of energy (emissions/energy), coupled with continuing increases in population and per-capita GDP. Nearly constant or slightly increasing trends in the carbon intensity of energy have been recently observed in both developed and developing regions. No region is decarbonizing its energy supply. The growth rate in emissions is strongest in rapidly developing economies, particularly China. Together, the developing and least-developed economies (forming 80% of the world's population) accounted for 73% of global emissions growth in 2004 but only 41% of global emissions and only 23% of global cumulative emissions since the mid-18th century. The results have implications for global equity.

Abstract

To better understand agricultural carbon fluxes in California, USA, we estimated changes in soil carbon and woody material between 1980 and 2000 on 3.6 x 10(6) ha of farmland in California. Combining the CASA (Carnegie-Ames-Stanford Approach) model with data on harvest indices and yields, we calculated net primary production, woody production in orchard and vineyard crops, and soil carbon. Over the 21-yr period, two trends resulted in carbon sequestration. Yields increased an average of 20%, corresponding to greater plant biomass and more carbon returned to the soils. Also, orchards and vineyards increased in area from 0.7 x 10(6) ha to 1.0 x 10(6) ha, displacing field crops and sequestering woody carbon. Our model estimates that California's agriculture sequestered an average of 19 g C x m(-2) x yr(-1). Sequestration was lowest in non-rice annual cropland, which sequestered 9 g C x m(-2) x yr(-1) of soil carbon, and highest on land that switched from annual cropland to perennial cropland. Land that switched from annual crops to vineyards sequestered 68 g C x m(-2) x yr(-1), and land that switched from annual crops to orchards sequestered 85 g C x m(-2) x yr(-1). Rice fields, because of a reduction in field burning, sequestered 55 g C x m(-2) x yr(-1) in the 1990s. Over the 21 years, California's 3.6 x 10(6) ha of agricultural land sequestered 11.0 Tg C within soils and 3.5 Tg C in woody biomass, for a total of 14.5 Tg C statewide. This is equal to 0.7% of the state's total fossil fuel emissions over the same time period. If California's agriculture adopted conservation tillage, changed management of almond and walnut prunings, and used all of its orchard and vineyard waste wood in the biomass power plants in the state, California's agriculture could offset up to 1.6% of the fossil fuel emissions in the state.

Diverse responses of phenology to global changes in a grassland ecosystemPROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICACleland, E. E., Chiariello, N. R., Loarie, S. R., Mooney, H. A., Field, C. B.2006; 103 (37): 13740-13744

Abstract

Shifting plant phenology (i.e., timing of flowering and other developmental events) in recent decades establishes that species and ecosystems are already responding to global environmental change. Earlier flowering and an extended period of active plant growth across much of the northern hemisphere have been interpreted as responses to warming. However, several kinds of environmental change have the potential to influence the phenology of flowering and primary production. Here, we report shifts in phenology of flowering and canopy greenness (Normalized Difference Vegetation Index) in response to four experimentally simulated global changes: warming, elevated CO(2), nitrogen (N) deposition, and increased precipitation. Consistent with previous observations, warming accelerated both flowering and greening of the canopy, but phenological responses to the other global change treatments were diverse. Elevated CO(2) and N addition delayed flowering in grasses, but slightly accelerated flowering in forbs. The opposing responses of these two important functional groups decreased their phenological complementarity and potentially increased competition for limiting soil resources. At the ecosystem level, timing of canopy greenness mirrored the flowering phenology of the grasses, which dominate primary production in this system. Elevated CO(2) delayed greening, whereas N addition dampened the acceleration of greening caused by warming. Increased precipitation had no consistent impacts on phenology. This diversity of phenological changes, between plant functional groups and in response to multiple environmental changes, helps explain the diversity in large-scale observations and indicates that changing temperature is only one of several factors reshaping the seasonality of ecosystem processes.

Abstract

In this study, the influence of elevated carbon dioxide (CO2) and nitrogen (N) deposition on gastropod herbivory was investigated for six annual species in a California annual grassland community. These experimentally simulated global changes increased availability of important resources for plant growth, leading to the hypothesis that species with the most positive growth and foliar nutrient responses would experience the greatest increase in herbivory. Counter to the expectations, shifts in tissue N and growth rates caused by N deposition did not predict shifts in herbivore consumption rates. N deposition increased seedling N concentrations and growth rates but did not increase herbivore consumption overall, or for any individual species. Elevated CO2 did not influence growth rates nor have a statistically significant influence on seedling N concentrations. Elevated CO2 at ambient N levels caused a decline in the number of seedlings consumed, but the interaction between CO2 and N addition differed among species. The results of this study indicate that shifting patterns of herbivory will likely influence species composition as environmental conditions change in the future; however, a simple trade-off between shifting growth rates and palatability is not evident.

Abstract

Selective consumption by herbivores influences the composition and structure of a range of plant communities. Anthropogenically driven global environmental changes, including increased atmospheric carbon dioxide (CO(2)), warming, increased precipitation, and increased N deposition, directly alter plant physiological properties, which may in turn modify herbivore consumption patterns. In this study, we tested the hypothesis that responses of annual grassland composition to global changes can be predicted exclusively from environmentally induced changes in the consumption patterns of a group of widespread herbivores, the terrestrial gastropods. This was done by: (1) assessing gastropod impacts on grassland composition under ambient conditions; (2) quantifying environmentally induced changes in gastropod feeding behaviour; (3) predicting how grassland composition would respond to global-change manipulations if influenced only by herbivore consumption preferences; and (4) comparing these predictions to observed responses of grassland community composition to simulated global changes. Gastropod herbivores consume nearly half of aboveground production in this system. Global changes induced species-specific changes in plant leaf characteristics, leading gastropods to alter the relative amounts of different plant types consumed. These changes in gastropod feeding preferences consistently explained global-change-induced responses of functional group abundance in an intact annual grassland exposed to simulated future environments. For four of the five global change scenarios, gastropod impacts explained > 50% of the quantitative changes, indicating that herbivore preferences can be a major driver of plant community responses to global changes.

Abstract

In this century, increasing concentrations of carbon dioxide (CO2) and other greenhouse gases in the Earth's atmosphere are expected to cause warmer surface temperatures and changes in precipitation patterns. At the same time, reactive nitrogen is entering natural systems at unprecedented rates. These global environmental changes have consequences for the functioning of natural ecosystems, and responses of these systems may feed back to affect climate and atmospheric composition. Here, we report plant growth responses of an ecosystem exposed to factorial combinations of four expected global environmental changes. We exposed California grassland to elevated CO2, temperature, precipitation, and nitrogen deposition for five years. Root and shoot production did not respond to elevated CO2 or modest warming. Supplemental precipitation led to increases in shoot production and offsetting decreases in root production. Supplemental nitrate deposition increased total production by an average of 26%, primarily by stimulating shoot growth. Interactions among the main treatments were rare. Together, these results suggest that production in this grassland will respond minimally to changes in CO2 and winter precipitation, and to small amounts of warming. Increased nitrate deposition would have stronger effects on the grassland. Aside from this nitrate response, expectations that a changing atmosphere and climate would promote carbon storage by increasing plant growth appear unlikely to be realized in this system.

Abstract

Although global changes can alter ecosystem nutrient dynamics indirectly as a result of their effects on plant litter quality, the interactive effects of global changes on plant litter remain largely unexplored in natural communities. We investigated the effects of elevated CO2, N deposition, warming and increased precipitation on the composition of organic compounds in plant litter in a fully-factorial experiment conducted in a California annual grassland. While lignin increased within functional groups under elevated CO2, this effect was attenuated by warming in grasses and by water additions in forbs. CO2-induced increases in lignin within functional groups also were counteracted by an increase in the relative biomass of forbs, which contained less lignin than grasses. Consequently, there was no net change in the overall lignin content of senesced tissue at the plot level under elevated CO2. Nitrate additions increased N in both grass and forb litter, although this effect was attenuated by water additions. Relative to changes in N within functional groups, changes in functional group dominance had a minor effect on overall litter N at the plot level. Nitrate additions had the strongest effect on decomposition, increasing lignin losses from Avena litter and interacting with water additions to increase decomposition of litter of other grasses. Increases in lignin that resulted from elevated CO2 had no effect on decomposition but elevated CO2 increased N losses from Avena litter. Overall, the interactions among elements of global change were as important as single-factor effects in influencing plant litter chemistry. However, with the exception of variation in N, litter quality had little influence on decomposition over the short term.

Ammonia-oxidizing bacteria respond to multifactorial global changePROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICAHorz, H. P., Barbrook, A., Field, C. B., Bohannan, B. J.2004; 101 (42): 15136-15141

Abstract

Recent studies have demonstrated that multiple co-occurring global changes can alter the abundance, diversity, and productivity of plant communities. Below ground processes, often mediated by soil microorganisms, are central to the response of these communities to global change. Very little is known, however, about the effects of multiple global changes on microbial communities. We examined the response of ammonia-oxidizing bacteria (AOB), microorganisms that mediate the transformation of ammonium into nitrite, to simultaneous increases in atmospheric CO2, precipitation, temperature, and nitrogen deposition, manipulated on the ecosystem level in a California grassland. Both the community structure and abundance of AOB responded to these simulated global changes. Increased nitrogen deposition significantly altered the structure of the ammonia-oxidizing community, consistently shifting the community toward dominance by bacteria most closely related to Nitrosospira sp. 2. This shift was most pronounced when temperature and precipitation were not increased. Total abundance of AOB significantly decreased in response to increased atmospheric CO2. This decrease was most pronounced when precipitation was also increased. Shifts in community composition were associated with increases in nitrification, but changes in abundance were not. These results demonstrate that microbial communities can be consistently altered by global changes and that these changes can have implications for ecosystem function.

Abstract

The magnitude of future climate change depends substantially on the greenhouse gas emission pathways we choose. Here we explore the implications of the highest and lowest Intergovernmental Panel on Climate Change emissions pathways for climate change and associated impacts in California. Based on climate projections from two state-of-the-art climate models with low and medium sensitivity (Parallel Climate Model and Hadley Centre Climate Model, version 3, respectively), we find that annual temperature increases nearly double from the lower B1 to the higher A1fi emissions scenario before 2100. Three of four simulations also show greater increases in summer temperatures as compared with winter. Extreme heat and the associated impacts on a range of temperature-sensitive sectors are substantially greater under the higher emissions scenario, with some interscenario differences apparent before midcentury. By the end of the century under the B1 scenario, heatwaves and extreme heat in Los Angeles quadruple in frequency while heat-related mortality increases two to three times; alpine/subalpine forests are reduced by 50-75%; and Sierra snowpack is reduced 30-70%. Under A1fi, heatwaves in Los Angeles are six to eight times more frequent, with heat-related excess mortality increasing five to seven times; alpine/subalpine forests are reduced by 75-90%; and snowpack declines 73-90%, with cascading impacts on runoff and streamflow that, combined with projected modest declines in winter precipitation, could fundamentally disrupt California's water rights system. Although interscenario differences in climate impacts and costs of adaptation emerge mainly in the second half of the century, they are strongly dependent on emissions from preceding decades.

Abstract

Models predict that global warming may increase aridity in water-limited ecosystems by accelerating evapotranspiration. We show that interactions between warming and the dominant biota in a grassland ecosystem produced the reverse effect. In a 2-year field experiment, simulated warming increased spring soil moisture by 5-10% under both ambient and elevated CO2. Warming also accelerated the decline of canopy greenness (normalized difference vegetation index) each spring by 11-17% by inducing earlier plant senescence. Lower transpirational water losses resulting from this earlier senescence provide a mechanism for the unexpected rise in soil moisture. Our findings illustrate the potential for organism-environment interactions to modify the direction as well as the magnitude of global change effects on ecosystem functioning.

Abstract

Biodiversity responses to ongoing climate and atmospheric changes will affect both ecosystem processes and the delivery of ecosystem goods and services. Combined effects of co-occurring global changes on diversity, however, are poorly understood. We examined plant diversity responses in a California annual grassland to manipulations of four global environmental changes, singly and in combination: elevated CO2, warming, precipitation, and nitrogen deposition. After 3 years, elevated CO2 and nitrogen deposition each reduced plant diversity, whereas elevated precipitation increased it and warming had no significant effect. Diversity responses to both single and combined global change treatments were driven overwhelmingly by gains and losses of forb species, which make up most of the native plant diversity in California grasslands. Diversity responses across treatments also showed no consistent relationship to net primary production responses, illustrating that the diversity effects of these environmental changes could not be explained simply by changes in productivity. In two- to four-way combinations, simulated global changes did not interact in any of their effects on diversity. Our results show that climate and atmospheric changes can rapidly alter biological diversity, with combined effects that, at least in some settings, are simple, additive combinations of single-factor effects.

Abstract

Simulated global changes, including warming, increased precipitation, and nitrogen deposition, alone and in concert, increased net primary production (NPP) in the third year of ecosystem-scale manipulations in a California annual grassland. Elevated carbon dioxide also increased NPP, but only as a single-factor treatment. Across all multifactor manipulations, elevated carbon dioxide suppressed root allocation, decreasing the positive effects of increased temperature, precipitation, and nitrogen deposition on NPP. The NPP responses to interacting global changes differed greatly from simple combinations of single-factor responses. These findings indicate the importance of a multifactor experimental approach to understanding ecosystem responses to global change.

Carbon emissions from tropical deforestation and regrowth based on satellite observations for the 1980s and 1990sPROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICADeFries, R. S., Houghton, R. A., Hansen, M. C., Field, C. B., Skole, D., Townshend, J.2002; 99 (22): 14256-14261

Abstract

Carbon fluxes from tropical deforestation and regrowth are highly uncertain components of the contemporary carbon budget, due in part to the lack of spatially explicit and consistent information on changes in forest area. We estimate fluxes for the 1980s and 1990s using subpixel estimates of percent tree cover derived from coarse (National Oceanic and Atmospheric Administration's Advanced Very High Resolution Radiometer) satellite data in combination with a terrestrial carbon model. The satellite-derived estimates of change in forest area are lower than national reports and remote-sensing surveys from the United Nations Food and Agriculture Organization Forest Resource Assessment (FRA) in all tropical regions, especially for the 1980s. However, our results indicate that the net rate of tropical forest clearing increased approximately 10% from the 1980s to 1990s, most notably in southeast Asia, in contrast to an 11% reduction reported by the FRA. We estimate net mean annual carbon fluxes from tropical deforestation and regrowth to average 0.6 (0.3-0.8) and 0.9 (0.5-1.4) petagrams (Pg).yr(-1) for the 1980s and 1990s, respectively. Compared with previous estimates of 1.9 (0.6-2.5) Pg.yr(-1) based on FRA national statistics of changes in forest area, this alternative estimate suggests less "missing" carbon from the global carbon budget but increasing emissions from tropical land-use change.

Abstract

Knowledge of carbon exchange between the atmosphere, land and the oceans is important, given that the terrestrial and marine environments are currently absorbing about half of the carbon dioxide that is emitted by fossil-fuel combustion. This carbon uptake is therefore limiting the extent of atmospheric and climatic change, but its long-term nature remains uncertain. Here we provide an overview of the current state of knowledge of global and regional patterns of carbon exchange by terrestrial ecosystems. Atmospheric carbon dioxide and oxygen data confirm that the terrestrial biosphere was largely neutral with respect to net carbon exchange during the 1980s, but became a net carbon sink in the 1990s. This recent sink can be largely attributed to northern extratropical areas, and is roughly split between North America and Eurasia. Tropical land areas, however, were approximately in balance with respect to carbon exchange, implying a carbon sink that offset emissions due to tropical deforestation. The evolution of the terrestrial carbon sink is largely the result of changes in land use over time, such as regrowth on abandoned agricultural land and fire prevention, in addition to responses to environmental changes, such as longer growing seasons, and fertilization by carbon dioxide and nitrogen. Nevertheless, there remain considerable uncertainties as to the magnitude of the sink in different regions and the contribution of different processes.

Abstract

For the period 1980-89, we estimate a carbon sink in the coterminous United States between 0.30 and 0.58 petagrams of carbon per year (petagrams of carbon = 10(15) grams of carbon). The net carbon flux from the atmosphere to the land was higher, 0.37 to 0.71 petagrams of carbon per year, because a net flux of 0.07 to 0.13 petagrams of carbon per year was exported by rivers and commerce and returned to the atmosphere elsewhere. These land-based estimates are larger than those from previous studies (0.08 to 0.35 petagrams of carbon per year) because of the inclusion of additional processes and revised estimates of some component fluxes. Although component estimates are uncertain, about one-half of the total is outside the forest sector. We also estimated the sink using atmospheric models and the atmospheric concentration of carbon dioxide (the tracer-transport inversion method). The range of results from the atmosphere-based inversions contains the land-based estimates. Atmosphere- and land-based estimates are thus consistent, within the large ranges of uncertainty for both methods. Atmosphere-based results for 1980-89 are similar to those for 1985-89 and 1990-94, indicating a relatively stable U.S. sink throughout the period.

Abstract

The Sea-viewing Wide Field-of-view Sensor (SeaWiFS) provides global monthly measurements of both oceanic phytoplankton chlorophyll biomass and light harvesting by land plants. These measurements allowed the comparison of simultaneous ocean and land net primary production (NPP) responses to a major El Niño to La Niña transition. Between September 1997 and August 2000, biospheric NPP varied by 6 petagrams of carbon per year (from 111 to 117 petagrams of carbon per year). Increases in ocean NPP were pronounced in tropical regions where El Niño-Southern Oscillation (ENSO) impacts on upwelling and nutrient availability were greatest. Globally, land NPP did not exhibit a clear ENSO response, although regional changes were substantial.

Abstract

Carbon accumulation in the terrestrial biosphere could partially offset the effects of anthropogenic CO2 emissions on atmospheric CO2. The net impact of increased CO2 on the carbon balance of terrestrial ecosystems is unclear, however, because elevated CO2 effects on carbon input to soils and plant use of water and nutrients often have contrasting effects on microbial processes. Here we show suppression of microbial decomposition in an annual grassland after continuous exposure to increased CO2 for five growing seasons. The increased CO2 enhanced plant nitrogen uptake, microbial biomass carbon, and available carbon for microbes. But it reduced available soil nitrogen, exacerbated nitrogen constraints on microbes, and reduced microbial respiration per unit biomass. These results indicate that increased CO2 can alter the interaction between plants and microbes in favour of plant utilization of nitrogen, thereby slowing microbial decomposition and increasing ecosystem carbon accumulation.

Abstract

Using a simple light-use efficiency model based on optical measurements, we explored spatial patterns of photosynthetic activity in fertilized and unfertilized sunflower stands. The model had two components: (1) absorbed photosynthetically active radiation (APAR), and (2) radiation-use efficiency. APAR was the product of photosynthetic photon flux density (PPFD) and leaf absorptance, which was derived from leaf reflectance. Radiation-use efficiency was either assumed to be constant or allowed to vary linearly with the photochemical reflectance index (PRI), a measure of xanthophyll cycle pigment activity. When efficiency was assumed to be constant, the model overestimated photosynthetic rates in upper canopy layers exposed to direct PPFD, particularly in the unfertilized canopy due to the greater photosynthetic downregulation associated with higher levels of photoprotective (de-epoxidized) xanthophyll cycle pigments in these conditions. When efficiency was allowed to vary according to the PRI, modeled photosynthetic rates closely matched measured rates for all canopy layers in both treatments. These results illustrate the importance of considering reduced radiation-use efficiency due to photosynthetic downregulation when modeling photosynthesis from reflectance, and illustrate the potential for detecting radiation-use efficiency through leaf optical properties. At least under the conditions of this study, these results also suggest that xanthophyll cycle pigment activity and net carbon uptake are coordinately regulated, allowing assays of Photosystem II activity to reveal changing rates of net assimilation. Because the optical methods in this study are adaptable to multiple spatial scales (leaf to landscape), this approach may provide a scalable model for estimating photosynthetic rates independently from flux measurements.

Abstract

In a seminal paper, Garrett Hardin argued in 1968 that users of a commons are caught in an inevitable process that leads to the destruction of the resources on which they depend. This article discusses new insights about such problems and the conditions most likely to favor sustainable uses of common-pool resources. Some of the most difficult challenges concern the management of large-scale resources that depend on international cooperation, such as fresh water in international basins or large marine ecosystems. Institutional diversity may be as important as biological diversity for our long-term survival.

Abstract

Integrating conceptually similar models of the growth of marine and terrestrial primary producers yielded an estimated global net primary production (NPP) of 104.9 petagrams of carbon per year, with roughly equal contributions from land and oceans. Approaches based on satellite indices of absorbed solar radiation indicate marked heterogeneity in NPP for both land and oceans, reflecting the influence of physical and ecological processes. The spatial and temporal distributions of ocean NPP are consistent with primary limitation by light, nutrients, and temperature. On land, water limitation imposes additional constraints. On land and ocean, progressive changes in NPP can result in altered carbon storage, although contrasts in mechanisms of carbon storage and rates of organic matter turnover result in a range of relations between carbon storage and changes in NPP.

Abstract

Atmospheric general circulation models used for climate simulation and weather forecasting require the fluxes of radiation, heat, water vapor, and momentum across the land-atmosphere interface to be specified. These fluxes are calculated by submodels called land surface parameterizations. Over the last 20 years, these parameterizations have evolved from simple, unrealistic schemes into credible representations of the global soil-vegetation-atmosphere transfer system as advances in plant physiological and hydrological research, advances in satellite data interpretation, and the results of large-scale field experiments have been exploited. Some modern schemes incorporate biogeochemical and ecological knowledge and, when coupled with advanced climate and ocean models, will be capable of modeling the biological and physical responses of the Earth system to global change, for example, increasing atmospheric carbon dioxide.

Abstract

Circadian rhythms in stomatal opening and photosynthesis had shorter free-running periods than circadian rhythms in leaflet movement in bean plants (Phaseolus vulgaris L.) transferred from 12-hr photoperiods to constant conditions. The rhythm in leaflet movement had a period close to 27 hr, whereas the rhythm in stomatal opening, measured as conductance to water vapor, had a period close to 24 hr. Photosynthesis, measured as net assimilation of CO2, also oscillated with a period close to 24 hr. The periods of these rhythms did not vary with increasing temperature, demonstrating temperature compensation of the controlling oscillators. The difference in free-running periods displayed by these rhythms is evidence that multiple oscillators with different intrinsic frequencies operate in bean plants.

Abstract

A circadian rhythm in photosynthesis occurs in Phaseolus vulgaris after transfer from a natural or artificial light:dark cycle to constant light. The rhythm in photosynthesis persists even when intercellular CO(2) partial pressure is held constant, demonstrating that the rhythm in photosynthesis is not entirely due to stomatal control over the diffusion of CO(2). Experiments were conducted to attempt to elucidate biochemical correlates with the circadian rhythm in photosynthesis. Plants were entrained to a 12-hour-day:12-hour-night light regimen and then monitored or sampled during a subsequent period of constant light. We observed circadian oscillations in ribulose-1,5-bisphosphate (RuBP) levels, and to a lesser extent in phosphoglyceric acid (PGA) levels, that closely paralleled oscillations in photosynthesis. However, the enzyme activity and activation state of the enzyme responsible for the conversion of RuBP to PGA, ribulose-1,5-bisphosphate carboxylase/oxygenase, showed no discernible circadian oscillation. Hence, we examined the possibility of circadian effects on RuBP regeneration. Neither ribulose-5-phosphate kinase activity nor the level of ATP fluctuated in constant light. Oscillations in triose-phosphate levels were out of phase with those observed for RuBP and PGA.

Abstract

Net carbon assimilation and stomatal conductance to water vapor oscillated repeatedly in red kidney bean, Phaseolus vulgaris L., plants transferred from a natural photoperiod to constant light. In a gas exchange system with automatic regulation of selected environmental and physiological variables, assimilation and conductance oscillated with a free-running period of approximately 24.5 hours. The rhythms in carbon assimilation and stomatal conductance were closely coupled and persisted for more than a week under constant conditions. A rhythm in assimilation occurred when either ambient or intercellular CO(2) partial pressure was held constant, demonstrating that the rhythm in assimilation was not entirely the result of stomatal effects on CO(2) diffusion. Rhythms in assimilation and conductance were not expressed in plants grown under constant light at a constant temperature, demonstrating that the rhythms did not occur spontaneously but were induced by an external stimulus. In plants grown under constant light with a temperature cycle, a rhythm was entrained in stomatal conductance but not in carbon assimilation, indicating that the oscillators driving the rhythms differed in their sensitivity to environmental stimuli.

Abstract

Many studies have shown that membrane lipids of chilling-sensitive plants begin lateral phase separation (i.e. a minor component begins freezing) at chilling temperatures and that chilling-sensitive plants are often of tropical origin. We tested the hypothesis that membranes of tropical plants begin lateral phase separation at chilling temperatures, and that plants lower the temperature of lateral phase separation as they invade cooler habitats. To do so we studied plant species in one family confined to the tropics (Piperaceae) and in three families with both tropical and temperate representatives (Fabaceae [Leguminosae], Malvaceae, and Solanaceae). We determined lateral phase separation temperatures by measuring the temperature dependence of fluorescence from trans-parinaric acid inserted into liposomes prepared from isolated membrane phospholipids. In all families we detected lateral phase separations at significantly higher temperatures, on average, in species of tropical origin. To test for associated physiological effects we measured the temperature dependence of delayed light emission (DLE) by discs cut from the same leaves used for lipid analysis. We found that the temperature of maximum DLE upon chilling was strongly correlated with lateral phase separation temperatures, but was on average approximately 4 degrees C lower. We also tested the hypothesis that photosystem II (PSII) (the most thermolabile component of photosynthesis) of tropical plants tolerates higher temperatures than PSII of temperate plants, using DLE and F(o) chlorophyll fluorescence upon heating to measure the temperature at which PSII thermally denatured. We found little difference between the two groups in PSII denaturation temperature. We also found that the temperature of maximum DLA upon heating was not significantly different from the critical temperature for F(o) fluorescence. Our results indicate that plants lowered their membrane freezing temperatures as they radiated from their tropical origins. One interpretation is that the tendency for membranes to begin freezing at chilling temperatures is the primitive condition, which plants corrected as they invaded colder habitats. An alternative is that membranes which freeze at temperatures only slightly lower than the minimum growth temperature confer an advantage.

ENVIRONMENTAL CONTROLS ON STOMATAL CONDUCTANCE IN A SHRUB OF THE HUMID TROPICSPROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA-BIOLOGICAL SCIENCESMooney, H. A., Field, C., YANES, C. V., Chu, C.1983; 80 (5): 1295-1297

Abstract

Leaves of Piper hispidum, a shrub native to the lowland tropics of Mexico, have a strong stomatal response to humidity that results in similar rates of water loss under a wide range of leaf-to-air water-vapor concentration gradients. Stomatal conductance of these leaves is insensitive to CO(2) concentration and increases in response to high humidity even in the dark.

Abstract

The photocontrol of the functional coupling between photosynthesis and stomatal conductance in the leaf was investigated in gas exchange experiments using monochromatic light provided by lasers. Net photosynthesis and stomatal conductance were measured in attached leaves of Malva parviflora L. as a function of photon irradiance at 457.9 and 640.0 nanometers.Photosynthetic rates and quantum yields of photosynthesis were higher under red light than under blue, on an absorbed or incident basis.Stomatal conductance was higher under blue than under red light at all intensities. Based on a calculated apparent photon efficiency of conductance, blue and red light had similar effects on conductance at intensities higher than 0.02 millimoles per square meter per second, but blue light was several-fold more efficient at very low photon irradiances. Red light had no effect on conductance at photon irradiances below 0.02 millimoles per square meter per second. These observations support the hypothesis that stomatal conductance is modulated by two photosystems: a blue light-dependent one, driving stomatal opening at low light intensities and a photosynthetically active radiation (PAR)-dependent one operating at higher irradiances.When low intensity blue light was used to illuminate a leaf already irradiated with high intensity, 640 nanometers light, the leaf exhibited substantial increases in stomatal conductance. Net photosynthesis changed only slightly. Additional far-red light increased net photosynthesis without affecting stomatal conductance. These observations indicate that under conditions where the PAR-dependent system is driven by high intensity red light, the blue light-dependent system has an additive effect on stomatal conductance.The wavelength dependence of photosynthesis and stomatal conductance demonstrates that these processes are not obligatorily coupled and can be controlled by light, independent of prevailing levels of intercellular CO(2). The blue light-dependent system in the guard cells may function as a specific light sensor while the PAR-dependent system supplies a CO(2)-modulated energy source providing functional coupling between the guard cells and the photosynthesizing mesophyll.